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Background and AimsDespite general agreement regarding the adaptive importance of plasticity, evidence for the role of environmental resource availability in plants is scarce. In arid and semi-arid environments, the persistence and dominance of perennial species depends on their capacity to tolerate drought: tolerance could be given on one extreme by fixed traits and, on the other, by plastic traits. To understand drought tolerance of species it is necessary to know the plasticity of their water economy-related traits, i.e. the position in the fixed–plastic continuum.MethodsThree conspicuous co-existing perennial grasses from a Patagonian steppe were grown under controlled conditions with four levels of steady-state water availability. Evaluated traits were divided into two groups. The first was associated with potential plant performance and correlated with fitness, and included above-ground biomass, total biomass, tillering and tiller density at harvest. The second group consisted of traits associated with mechanisms of plant adjustment to environmental changes and included root biomass, shoot/root ratio, tiller biomass, length of total elongated leaf, length of yellow tissue divided by time and final length divided by the time taken to reach final length.Key Results and ConclusionsThe most plastic species along this drought gradient was the most sensitive to drought, whereas the least plastic and slowest growing was the most tolerant. This negative relationship between tolerance and plasticity was true for fitness-related traits but was trait-dependent for underlying traits. Remarkably, the most tolerant species had the highest positive plasticity (i.e. opposite to the default response to stress) in an underlying trait, directly explaining its drought resistance: it increased absolute root biomass. The niche differentiation axis that allows the coexistence of species in this group of perennial dryland grasses, all limited by soil surface moisture, would be a functional one of fixed versus plastic responses.

Nutrient limitation is a basic ecological constraint that has received little attention in studies on virus production and disease dynamics. Nutrient availability could directly limit the production of viral nucleic acids and proteins, or alternatively limit host growth and thus indirectly limit metabolic pathways necessary for viral replication. In order to compare direct and indirect effects of nutrient limitation on virus production within hosts, we manipulated soil nitrogen (N) and phosphorus (P) availability in a glasshouse for the wild grass host Bromus hordeaceus and the viral pathogen Barley yellow dwarf virus-PAV. We found that soil N additions increased viral concentrations within host tissues, and the effect was mediated by host growth. Specifically, in statistical models evaluating the roles of host biomass production, leaf N and leaf P, viral production depended most strongly on host biomass, rather than the concentration of either nutrient. Furthermore, at low soil N, larger plants supported greater viral concentrations than smaller ones, whereas at high N, smaller plants supported greater viral concentrations. Our results suggest that enhanced viral productivity under N enrichment is an indirect consequence of nutrient stimulation to host growth rate. Heightened pathogen production in plants has important implications for a world facing increasing rates of nutrient deposition.

Exotic invasive species can directly and indirectly influence natural ecological communities. Cheatgrass (Bromus tectorum) is non-native to the western United States and has invaded large areas of the Great Basin. Changes to the structure and composition of plant communities invaded by cheatgrass likely have effects at higher trophic levels. As a keystone guild in North American deserts, granivorous small mammals drive and maintain plant diversity. Our objective was to assess potential effects of invasion by cheatgrass on small-mammal communities. We sampled small-mammal and plant communities at 70 sites (Great Basin, Utah). We assessed abundance and diversity of the small-mammal community, diversity of the plant community, and the percentage of cheatgrass cover and shrub species. Abundance and diversity of the small-mammal community decreased with increasing abundance of cheatgrass. Similarly, cover of cheatgrass remained a significant predictor of small-mammal abundance even after accounting for the loss of the shrub layer and plant diversity, suggesting that there are direct and indirect effects of cheatgrass. The change in the small-mammal communities associated with invasion of cheatgrass likely has effects through higher and lower trophic levels and has the potential to cause major changes in ecosystem structure and function.

Shifting precipitation patterns resulting from global climate change will influence the success of invasive plant species. In the Front Range of Colorado, Bromus tectorum (cheatgrass) and other non-native winter annuals have invaded grassland communities and are becoming more abundant. As the global climate warms, more precipitation may fall as rain rather than snow in winter, and an increase in winter rain could benefit early-growing winter annuals, such as B. tectorum, to the detriment of native species. In this study we measured the effects of simulated changes in seasonal precipitation and presence of other plant species on population growth of B. tectorum in a grassland ecosystem near Boulder, Colorado, USA. We also performed elasticity analyses to identify life transitions that were most sensitive to precipitation differences. In both study years, population growth rates were highest for B. tectorum growing in treatments receiving supplemental winter precipitation and lowest for those receiving the summer drought treatment. Survival of seedlings to flowering and seed production contributed most to population growth in all treatments. Biomass of neighboring native plants was positively correlated with reduced population growth rates of B. tectorum. However, exotic plant biomass had no effect on population growth rates. This study demonstrates how interacting effects of climate change and presence of native plants can influence the population growth of an invasive species. Overall, our results suggest that B. tectorum will become more invasive in grasslands if the seasonality of precipitation shifts towards wetter winters and allows B. tectorum to grow when competition from native species is low.

Climate change effects on plants are expected to be primarily mediated through early life stage transitions. Snowfall variability, in particular, may have profound impacts on seedling recruitment, structuring plant populations and communities, especially in mid-latitude systems. These water-limited and frequently invaded environments experience tremendous variation in snowfall, and species in these systems must contend with harsh winter conditions and frequent disturbance. In this study, we examined the mechanisms driving the effects of snowpack depth and soil disturbance on the germination, emergence, and establishment of the native Pseudoroegnaria spicata and the invasive Bromus tectorum, two grass species that are widely distributed across the cold deserts of North America. The absence of snow in winter exposed seeds to an increased frequency and intensity of freeze-thaw cycles and greater fungal pathogen infection. A shallower snowpack promoted the formation of a frozen surface crust, reducing the emergence of both species (more so for P. spicata). Conversely, a deeper snowpack recharged the soil and improved seedling establishment of both species by creating higher and more stable levels of soil moisture availability following spring thaw. Across several snow treatments, experimental disturbance served to decrease the cumulative survival of both species. Furthermore, we observed that, regardless of snowpack treatment, most seed mortality (70-80 %) occurred between seed germination and seedling emergence (November-March), suggesting that other wintertime factors or just winter conditions in general limited survival. Our results suggest that snowpack variation and legacy effects of the snowpack influence emergence and establishment but might not facilitate invasion of cold deserts.

Plant responses to changes in temperature can be a key factor in predicting the presence and managing invasive plant species while conserving resident native plant species in dryland ecosystems. Climate can influence germination, establishment, and seedling biomass of both native and invasive plant species. We tested the hypothesis that common and widely distributed native and an invasive plant species in dryland ecosystems in California respond differently to increasing temperatures. To test this, we examined the effects of temperature variation on germination, establishment, and per capita seedling biomass of three native and one invasive plant species ( Bromus rubens ) in independent 6 week growth trial experiments in a controlled greenhouse. Higher relative temperatures reduced the germination and establishment of the tested invasive species and two tested native species, however, per capita biomass was not significantly affected. Specifically, germination and establishment of the invasive species B. rubens and the native species Phacelia tanacetifolia was significantly reduced. This invasive species can often outcompete natives, but increasing temperature could potentially shift the balance between the germination and establishment of natives. A warming climate will likely have negative impacts on native annual plant species in California tested here because increasing temperatures can co-occur with drought. This study shows that our tested native annual plant species tested here have some resilience to relatively significant increases in temperature, and this can favor at least one native species relative to at least one highly noxious invasive plant species.

Climate change and disturbance can alter invasion success of clonal plants by differentially affecting the clonal traits influencing their establishment as young plants. Clonal traits related to the vegetative reproduction of native Pascopyrum smithii and non-native Bromus inermis grass seedlings were evaluated under altered precipitation frequencies and a single grazing event. Pascopyrum smithii maintained similar vegetative reproduction under three simulated precipitation frequencies whereas B. inermis vegetative reproduction declined as precipitation became more intermittent. Vegetative reproduction of the non-native B. inermis was greater than the native P. smithii under all simulated precipitation frequencies except the most intermittent scenario. A single grazing event did not affect either species’ response to intra-annual precipitation variability but did slightly reduce their clonal growth and increase their bud dormancy. In young plants, clonal traits of the invasive grass favored its superior expansion and population growth compared to the native grass except under the most severe climate change scenario. Grassland restoration using native P. smithii seeds would be successful in most years due to its resilient clonal growth in a changing climate. Clonal infrastructure development in young plants is critical to clonal plant establishment and persistence in a changing climate and under disturbed conditions.

Environmental extremes resulting from a changing climate can have profound implications for plant interactions in desert communities. Positive interactions can buffer plant communities from abiotic stress and consumer pressure caused by climatic extremes, but limited research has explored this empirically. We tested the hypothesis that the mechanism of shrub facilitation on an annual plant community can change with precipitation extremes in deserts. During years of extreme drought and above-average rainfall in a desert, we measured plant interactions and biomass while manipulating a soil moisture gradient and reducing consumer pressure. Shrubs facilitated the annual plant community at all levels of soil moisture through reductions in microclimatic stress in both years and herbivore protection in the wet year only. Shrub facilitation and the high rainfall year contributed to the dominance of a competitive annual species in the plant community. Precipitation patterns in deserts determine the magnitude and type of facilitation mechanisms. Moreover, shrub facilitation mediates the interspecific competition within the associated annual community between years with different rainfall amounts. Examining multiple drivers during extreme climate events is a challenging area of research, but it is a necessary consideration given forecasts predicting that these events will increase in frequency and magnitude.

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C₃ and C₄). The response of these communities was compared to an invaded/disturbed grassland. The invaded/disturbed communities had higher surface NH₄⁺ input in spring, whereas there were no differences for surface input of NO₃⁻. Soil inorganic N was dominated by NH₄⁺, but invaded sites had greater subsurface soil NO₃⁻. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil$\\delta {}^{15}\\text{N}$increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar$\\delta {}^{15}\\text{N}$values suggest that Bromus assimilated subsurface NO₃⁻, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil$\\delta {}^{15}\\text{N}$; however, the invaded/disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO₃⁻ concentrations were greater in the invaded communities during periods of ample precipitation, and NO₃⁻ leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO₃⁻ from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.

The possibility of increased invasiveness in cultivated varieties of native perennial species is a question of interest in biofuel risk assessment. Competitive success is a key factor in the fitness and invasive potential of perennial plants, and thus the large-scale release of high-yielding biomass cultivars warrants empirical comparisons with local conspecifics in the presence of competitors. We evaluated the performance of non-local cultivars and local wild biotypes of the tallgrass species Panicum virgatum L. (switchgrass) in competition experiments during two growing seasons in Ohio and Iowa. At each location, we measured growth and reproductive traits (plant height, tiller number, flowering time, aboveground biomass, and seed production) of four non-locally sourced cultivars and two locally collected wild biotypes. Plants were grown in common garden experiments under three types of competition, referred to as none, moderate (with Schizachyrium scoparium), and high (with Bromus inermis). In both states, the two \"lowland\" cultivars grew taller, flowered later, and produced between 2x and 7.5x more biomass and between 3x and 34x more seeds per plant than local wild biotypes, while the other two cultivars were comparable to wild biotypes in these traits. Competition did not affect relative differences among biotypes, with the exception of shoot number, which was more similar among biotypes under high competition. Insights into functional differences between cultivars and wild biotypes are crucial for developing biomass crops while mitigating the potential for invasiveness. Here, two of the four cultivars generally performed better than wild biotypes, indicating that these biotypes may pose more of a risk in terms of their ability to establish vigorous feral populations in new regions outside of their area of origin. Our results support an ongoing assessment of switchgrass cultivars developed for large-scale planting for biofuels.